UNIVERSITI PUTRA MALAYSIA

IMPACT OF SALINITY ON SEED GERMINATION AND ANTIOXIDATIVE PROFILE OF TOMATO LEAVES (Solanum lycopersicum L. cv.MT1)

ASMA ABUBAKER A. SHANAB

FS 2016 23

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IMPACT OF SALINITY ON SEED GERMINATION AND ANTIOXIDATIVE PROFILE OF TOMATO LEAVES (Solanum lycopersicum L. cv.MT1)

By

ASMA ABUBAKER A. SHANAB

Thesis Submitted to the School of Graduate Studies, Universiti Putra Malaysia, in Fulfilment of the Requirements for the Degree of

Master of Science

January 2016

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COPYRIGHT

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Abstract of thesis presented to the Senate of Universiti Putra Malaysia in fulfilment of the requirement for the Degree of Master of Science

IMPACT OF SALINITY ON SEED GERMINATION AND ANTIOXIDATIVE PROFILE OF TOMATO LEAVES (Solanum lycopersicum L. cv.MT1)

By

ASMA ABUBAKER A.SHANAB

January 2016

Chairman : Rosimah binti Nulit, PhD Faculty : Science Salinity is a serious problem in many parts of the world that limits the productivity of agricultural crops. Even though, it is one of the major abiotic stresses that affect almost every aspect of physiology and biochemistry of plants but at certain amount could be beneficial to plant. Many studies had been done to determine the effects of salt stress towards the morphology, physiology, biochemistry and productivity of tomato plant but study on germination and antioxidant activity on tomato plants is still limited. This work aimed to study the effects of NaCl on the germination, histological of seedling leaves, and antioxidant profiling of Solanum lycopersicum cv. MT1. A 20 surface sterilized of tomato MT1 seeds were germinated in sterilized petri dishes and treated with 6 ml of 1, 3, 5, 10, 15, 30 mM NaCl separately and deionized water used as a control to study the effects of NaCl on tomato seed germination. Plates were sealed with parafilm to prevent evaporation followed by placed in Random Block Design in the growth room at 25˚C and 16/8 h (light / dark) for 10 days. Seeds were considered germinated when radicle had extended at least 2 mm. The water uptake, germination percentage (% GP), seed vigor, salt tolerance, length of hypocotyl and radical, and biomass of seedlings were measured. Results found that the germination of MT1 seed adversely delayed and decreased as NaCl concentration increased. The water uptake, germination percentage, seed vigor and salt tolerance decreased when the concentration of NaCl increased except 3 mM NaCl showed the highest germination percentage (87.5%) and vigor (11.4). The water uptake, seed vigor and salt tolerance decreased when as the increased of NaCl. The response of hypocotyls and radical under NaCl treatment are different which is not significantly (ANOVA, P<0.05) affected the length of hypocotyl but significantly reduced the radical length (ANOVA, P<0.05) except in 3 mM NaCl. NaCl increase the seedling length up to 3 mM (13 cm). Increased NaCl concentration not significantly (ANOVA, P<0.05) reduced the shoot dry weight (ANOVA, P<0.05), on other hand, the root dry weight reduce significantly (ANOVA, P<0.05) as NaCl concentration higher. At day 10, seedling leaves were collected from control, 10 and 30 mM NaCl and fixed for histological studies. Histological of seedlings leaves highly affected by NaCl concentration. The entire structural of histological seedling leaves highly disrupted as NaCl

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concentration increased as can be compared between 10 mM and 30 mM NaCl. The structure of epidermis cells on both epidermal layers totally changed. Moreover, the arrangement of mesophylls tissues was also in disorder. A major reduction in veins tissue dimensions were observed and this lead on its ability to conduct water and nutrients into seedlings system. To study the effects of NaCl to the antioxidant profiling of tomato plant. Four different concentration of NaCl treatment which is 5, 10, 15 and 30 mM and deionized water as a control was used. 21 days old seedlings were planted in the pots containing 1 kg of mix soil (3 top soil: 1 sand). Each treatment has 6 replicates. The seedlings were watered with 100 ml of NaCl and alternate with 100 ml ultra-pure water every 3 days for six weeks. After 6 six weeks, total chlorophyll, total phenolic, flavonoid and proline and anti-oxidative activity were measured. Results showed that NaCl increase total chlorophyll, total phenolic, flavonoid and proline and anti-oxidative activity. Total chlorophyll found higher in 15 and 30 Mm NaCl which is 41.7 mg and 39.6 mg/0.1 g compared with control plant only 24.1 mg/0.1 g. Results showed an increasing of total phenolic, flavonoid, and proline content with increasing of NaCl. The highest amount of phenolic and flavonoid achieved at 15 mM NaCl higher than the control plant and other treatment. Results also found that total proline increase as NaCl increased. 15 and 30 mM NaCl show higher total proline content which is 0.041 and 0.044 mg/0.1g compared with control (0.016 mg/0.1g) and other NaCl treatment. The maximum of DPPH and reducing power activity were observed in the 5 mM NaCl while antioxidant reduced as NaCl increase up to 30 mM NaCl. As conclusion, raising the concentration of NaCl had adversely affected on the germination and early growth of seedlings (except 3 mM NaCl). The histological of MT1 seedlings leaves highly affected by NaCl. Treatment tomato plants with moderate concentration of NaCl able to increase the anti-oxidative properties and activity of tomato plant.

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Abstrak tesis yang dikemukakan kepada Senat Universiti Putra Malaysia sebagai memenuhi keperluan untuk Ijazah Master Sains

KESAN KEMASINAN KE ATAS PERCAMBAHAN BIJI BENIH DAN PROFIL ANTIOKSIDAN DAUN TOMATO (Solanum lycopersicum L. cv. MT1)

Oleh

ASMA ABUBAKER A.SHANAB

Januari 2016

Pengerusi : Rosimah binti Nulit, PhD Fakulti : Sains Kemasinan merupakan masalah serius di kebanyakan tempat di dunia dimana menghadkan produktiviti tanaman. Walaupun, kemasinan salah satu faktor abiotik yang memberi kesan kepada hampir setiap aspek fisiologi dan biokimia tumbuhan tetapi pada amaun tertentu bermenfaat kepada tumbuhan. Kesan tegasan kemasinan terhadap morfologi, fisiologi, biokimia dan produktiviti ke atas tomato telah banyak tetapi kajian ke atas percambahan dan aktiviti antioksidan pada tanaman tomato masih terhad. Kajian ini bertujuan untuk mengkaji kesan NaCl pada percambahan, histologi anatomi, dan profil antioksidan tomato (Solanum lycopersicum cv. MT1). Satu siri kepekatan NaCl iaitu 1, 3, 5, 10, 15, 30 mM dan air ternyahion sebagai kawalan telah digunakan untuk mengkaji kesan NaCl ke atas percambahan benih tomato. Sejumlah 20 biji benih tomato MT1 dimana permukaannya disteril terlebih dahulu disemai di dalam piring petri dan diberi rawatan dengan 6 ml larutan NaCl pada kepekatan 1, 3, 5, 10, 15, 30 mM NaCl dan air ternyahion digunakan sebagai kawalan. Semua piring petri dimeterai dengan parafilm untuk mengelakkan penyejatan dan ditempatkan secara blok rawak dalam bilik kultur pada suhu 25°C dan 16/8 h (terang/gelap) selama 10 hari. Biji benih dianggap bercambah apabila radikel telah tumbuh sekurang-kurangnya 2 mm. Peratusan percambahan (% GP), vigor bijibenih, toleransi terhadap kemasinan, panjang hipokotil dan radikel serta biomas anak benih diukur. Kajian mendapati, percambahan dan vigor biji benih MT1 tomato adalah lambat dan menurun dengan meningkatnya kepekatan NaCl kecuali pada rawatan 3 mM NaCl dimana peratus percambahan (87.5%) and vigor adalah yang tertinggi (11.4). Kajian juga mendapati gerakbalas hipokotil dan radikel terhadap NaCl berbeza dimana NaCl tidak memberikan kesan yang signifikan (ANOVA, P<0.05) ke atas kepanjangan hipokotil dan biomas pucuk tetapi mengurangkan pemanjangan radikel dan biomas akar (ANOVA, P<0.05) kecuali pada rawatan 3 mM NaCl. Pada hari ke sepuluh, daun anak-anak benih daripada kawalan, 10 mM dan 30 mM NaCl telah di awet untuk kajian histologi. Hasil kajian ini juga mendapati histologi daun anak benih sangat dipengaruhi oleh kepekatan NaCl. Keseluruhan struktur histologi daun anak benih tomato berubah apabila kepekatan NaCl meningkat seperti yang ditunjukkan pada kepekatan 10 mM dan 30 mM NaCl. Struktur dan bentuk sel-sel epidermis

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pada kedua-dua lapisan epidermis juga berubah. Kajian juga mendapati susunan kedua-dua jenis tisu mesofil tidak teratur. Di samping itu, pengurangan saiz tisu veina sangat ketara pada rawatan 10 mM dan 30 mM NaCl dan ini mengurangkan keupayaan pengangkutan air dan nutrient ke dalam anak benih. Untuk mengkaji kesan NaCl ke atas profil antioksidan pokok tomato, empat kepekatan NaCl iaitu 5, 10, 15 dan 30 mM dan air ternyahion sebagai kawalan digunakan. Anak benih yang berumur 21 hari ditanam di dalam pot yang mengandungi 1 kg tanah campuran (3 tanah atas:1 pasir). Setiap rawatan mengandungi 6 replikasi. Anak-anak benih disiram dengan 100 NaCl dan diselangi dengan 100 ml air ternyahion setiap 3 hari selama 6 minggu. Jumlah klorofil, jumlah fenolik, flavonoid, prolina dan aktiviti antioksidan diukur. Hasil kajian mendapati rawatan NaCl meningkatkan kandungan klorofil, fenolik, flavonoid dan prolina. Kandungan klorofil didapati tinggi pada rawatan 15 dan 30 mM NaCl iaitu 41.7 mg/0.1g dan 39.6 mg/0.1g berbanding tumbuhan kawalan hanya 24.1 mg/0.1g dan lain-lain rawatan. Jumlah tertinggi fenolik dan flavonoid ditunjukkan pada 15 mM NaCl. Hasil kajian mendapati kandungan prolina meningkat dengan peningkatan NaCl (ANOVA, p<0.05) dimana 15 dan 30 mM NaCl menunjukkan kandungan prolina 0.041 mg/0.1g dan 0.044 mg/0.1g berbanding kawalan hanya 0.016 mg/0.1g. Hasil kajian menunjukkan bahawa aktiviti DPPH dan kuasa penurunan adalah tertinggi dalam 5 mM NaCl dan semakin menurun pada kepekatan NaCl yang tinggi. Kesimpulannya, peningkatan kepekatan NaCl menyebabkan percambahan dan pertumbuhan awal anak benih lambat dan menurun (kecuali pada 3 mM NaCl). NaCl menjejaskan histologi daun anak benih tomato MT1. Rawatan garam dapat meningkatkan nilai antioksidan dan juga aktiviti pokok tomato MT1.

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ACKNOWLEDGEMENTS The name of Allah, first and foremost, all praise to Allah Subhanahu wa ta‘ala for bestowing me with health, opportunity, patience and knowledge to complete this thesis smoothly and the peace and blessings of Allah Subhanahu wa ta‘ala be upon Prophet Mohammed (peace be upon him). I would like to express my very great appreciation to my supervisor Dr Rosimah Nulit اfor helping, supporting and guiding me for the completion of this thesis as well for all the knowledge she had pass to me in all aspects of her guidance and supervision enabled me to complete my work successfully. I am indebted to her where, without her gaudiness this study could not be completed smoothly. I am also indebted to Dr. Mashitah Maidin my co-supervisor for her time and advice, as well as for her encouragement. My appreciation and honest thanks are due to all lecturers and staff members in department of Biology. I would like to offer my deep thanks to my husband Marwan for his continuous encouragement and in particular my beloved sons Abdul Malek and Abdul Aziz and my grateful thanks are also extended to my beloved parents and my parents in law for their continuous support and motivation, I pray to Allah to bless them, I would like to offer my deep thanks to my father for his continuous encouragement and helping me, also, I wish to express my sincere gratitude to my sister Sanaa and my brothers for their constant encouragement, love and ever ready to give helping hands. Finally yet importantly, special thanks to my friends for their generous support and encouragement in particular, Mabrokah, Nahid, Atiqah, Azia, Najlaa, Azimah and Mona and others whom not listed along.

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This thesis was submitted to the Senate of Universiti Putra Malaysia and has been accepted as fulfilment of the requirement for the degree of Master of Science. The members of the Supervisory Committee were as follows: Rosimah binti Nulit, PhD Senior Lecturer Faculty of Science Universiti Putra Malaysia (Chairman) Mashitah Binti Shikh Maidin, PhD Senior Lecturer Faculty of Science Universiti Putra Malaysia (Member)

BUJANG BIN KIM HUAT, PhD Professor and Dean School of Graduate Studies Universiti Putra Malaysia Date:

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Declaration by graduate student I hereby confirm that: this thesis is my original work; quotations, illustrations and citations have been duly referenced; this thesis has not been submitted previously or concurrently for any other

degree at any other institutions; intellectual property from the thesis and copyright of thesis are fully-owned

by Universiti Putra Malaysia, as according to the Universiti Putra Malaysia (Research) Rules 2012;

written permission must be obtained from supervisor and the office of Deputy Vice-Chancellor (Research and Innovation) before thesis is published (in the form of written, printed or in electronic form) including books, journals, modules, proceedings, popular writings, seminar papers, manuscripts, posters, reports, lecture notes, learning modules or any other materials as stated in the Universiti Putra Malaysia (Research) Rules 2012;

there is no plagiarism or data falsification/fabrication in the thesis, and scholarly integrity is upheld as according to the Universiti Putra Malaysia (Graduate Studies) Rules 2003 (Revision 2012-2013) and the Universiti Putra Malaysia (Research) Rules 2012. The thesis has undergone plagiarism detection software.

Signature: _______________________ Date: __________________ Name and Matric No.: Asma Abubaker A. Shanab, GS36547

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Declaration by Members of Supervisory Committee This is to confirm that: the research conducted and the writing of this thesis was under our

supervision; supervision responsibilities as stated in the Universiti Putra Malaysia

(Graduate Studies) Rules 2003 (Revision 2012-2013) are adhered to.

Signature: Name of Chairman of Supervisory Committee:

Rosimah binti Nulit, PhD

Signature:

Name of Member of Supervisory Committee:

Mashitah Binti Shikh Maidin, PhD

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TABLE OF CONTENTS

Page

ABSTRACT i ABSTRAK iii ACKNOWLEDGEMENTS v APPROVAL vi DECLARATION viii LIST OF TABLES xii LIST OF FIGURES xiii LIST OF APPENDICES xiv LIST OF ABBREVATIONS xv CHAPTER 1 INTRODUCTION 1.1 Background of study 1 1.2 Problem statements and objectives 2 2 LITERATURE REVIEW 2.1 Tomato 3 2.1.1 Introduction 3

2.1.2 Solanum lycopersicon L. cv. MT1 3 2.2 Salts in soils and plants 4 2.3 Salinity effects on the production of tomato 5 2.4 Effects of Salt Stress on Plants 6 2.5 Abiotic and biotic stress and their impact on plant

growth and productivity 7

2.6 Salinity Effects on seed germination 8 2.7 Effects of salinity on plant growth 9 2.8 Salinity effects on physiological, biochemical and anti-

oxidative profiling of plants 9

2.8.1 Physiological and biochemical changes as response to salinity

9

2.8.2 Anti-oxidative profiling affected by salinity 10 2.9 Salinity effects on the anatomy of plant 11 3 MATERIALS AND METHODS 3.1 Seed materials and seed sterilization 13 3.2 Preparation of sodium chloride (NaCl) solution 13 3.3 Experimental setup 13

3.3.1 Water uptake 13 3.3.2 The effect of salinity on germination 13

3.3.3 Measurement of growth of tomato seedlings 14 3.4 The effect of salinity on the histological leaves 14 3.4.1 Plant materials 14 3.4.2 Dehydration 14 3.4.3 Paraffin bath 15 3.4.4 Embedding and sectioning 15

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3.4.5 Staining 15 3.4.6 Slides mounting 15 3.5 The effects of NaCl on the tomato plant 15 3.5.1 Salinity treatment 15 3.6 Measurement of chlorophyll a, b and carotenoid 16 3.7 Measurement of total phenolic content 16 3.8 Measurement of flavonoid content 16 3.9 Measurement of proline content 16 3.10 Measurement of antioxidant activity assay 17 3.10.1 DPPH radical scavenging activity 17 3.10.2 Reducing power assay 18

3.11 Analysis of data 18

4 RESULTS 4.1 Water uptake percentage 19 4.2 Germination percentage 19

4.3 Seed vigor 21 4.4 Salt tolerance 21 4.5 Effects of salinity on early seedling growth 22 4.6 The effect of NaCl on histology of tomato seedling

leaves 24

4.7 The effects of NaCl on the photosynthetic pigments 26 4.8 The effects of NaCl on the total phenol, flavonoid and

proline 27

4.9 The antioxidant activity as respond to NaCl 28 4.9.1 2, 2- diphenyl- 1 picrylhydrazyl (DPPH)

Radical-scavenging activity assay 28

4.9.2 Reducing power assay 30 5 DISCUSSION

5.1 Effect of NaCl on germination of tomato seed 32 5.2 Effect of NaCl on the histological of MT1 seedling

leaves 34

5.3 Effect of salinity on anti-oxidative profiling of MT1 tomato plants

34

6 CONCLUSION AND RECOMMENDATION Conclusion 38 Recommendation 38 REFERENCES 39 APPENDICES 56 BIODATE OF STUDENT 73

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LIST OF TABLES

Table

Page

4.1 Effects of NaCl on the water uptake and seed germination of tomato seed

20

4.2 Effects of different levels of salt stress on seed vigor and salt tolerance of tomato

21

4.3 Effects of NaCl on hypocotyl length, radical length, and seedling length of tomato seedlings

23

4.4 Effects of NaCl on shoot and root dry weights of tomato seedlings

24

4.5 Chlorophyll a, chlorophyll b, total chlorophyll and carotenoid on week 6 in tomato plants as respond to different concentration of NaCl

27

4.6 The contents of phenolic, flavonoid, and proline of tomato leaves under different NaCl concentration

28

4.7 IC50 values of MT1 tomato leaves as respond to salinity

29

4.8

Comparison of Reducing Power between different Concentrations of NaCl of tomato leaf extracts at concentration of 10 mg/ml samples

31

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LIST OF FIGURES

Figure

Page

2.1 Categorization of the world soils in terms of salinity

6

4.1 Germination of tomato seeds under different concentration of NaCl on day 10

20

4.2 Lengths of hypocotyls and radicles of tomato seedlings germinated in NaCl after 10 days of treatment

22

4.3a Cross-sections of the leaves of tomato seedlings in control

25

4.3b Cross-sections of the leaves of tomato seedlings in 10 mM NaCl

25

4.3c Cross-sections of the leaves of tomato seedlings in 30 mM NaCl

26

4.4 DPPH radical scavenging activity of tomato leaves extract under different concentration of NaCl

29

4.5 The reducing power assay of tomato leaf extracts under different concentrations of NaCl

30

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LIST OF APPENDICES

Appendix

Page

1 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons of Water Uptake.

56

2 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple Comparisons of Germination Percentage.

57

3 One way ANOVA, Tukey HSD and Duncan multiple comparison for Seed Vigor

58

4 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons for Salt Tolerance.

59

5 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons for Shoot, Root and Seedling Length

60

6 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons for Shoot and Root Dry Weight.

63

7 One way ANOVA, Tukey HSD, Tukey B and Duncan Multiple Comparisons for Chlorophyll a, b, total Chlorophyll and Carotenoid.

65

8 One way ANOVA, Tukey HSD, Tukey B and Duncan Multiple comparisons of Flavonoids and Phenolic.

68

9 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons of Proline.

70

10 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons of IC50

71

11 One way ANOVA, Tukey HSD, Tukey B and Duncan multiple comparisons of Reducing Power Assay

72

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LIST OF ABBREVATIONS

% percentage ˚C degree Celsius µl Microliter AlCl3 Aluminium chloride ANOVA Analysis of Variance Ca2+ Calcium ion Cl- Chloride ion Cm Centimetre CO2 Carbon Dioxide Cu Copper DPPH 2, 2- diphenyl-1-picrylhidrazy DPX Diputal petroleum xylene DW Dry weight Fe Iron FW Fresh Weight G Gram Mg Milligram Mg2+ magnesium ion MgCl2 Magnesium chloride mg/µl milligram per microliter mg/l milligram per liter ml Milliliter mM miliMolar Na+ Ammonia Na2co3 Sodium carbonate NaCl Sodium Chloride NADH nicotinamide adenine dinucleotide reduced Nm Nanometre PSI Photosystem I ROS Reactive Oxygen Species v/v volume per volume w/v weight per volume

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CHAPTER 1

INTRODUCTION 1.1 Background of study Nowadays, the phenomenon of the salinity is one of the most serious problems facing soil and also effecting seed germination and plant growth. It is becoming one of the issues that hamper the economy in most countries of the world. Salinity is a common term used to describe the presence of elevated levels of different salt content such as sodium chloride, magnesium and calcium sulphates in soil and water. It has become a major problem in agriculture as high concentration of minerals in soil that causes salt stress to plant. Salinization of soil is one of the major factors limiting crop reproduction particularly in semi-arid and arid regions of the world (Ahmed, 2009). Salt stress can be defined as excess in ions especially in sodium ions (Na+) and chloride ions (Cl-). This salt stress reduces the ability of the plant to take up water which can reduce the growth rate as well as causes metabolic changes similar to water stress conditions (Munns, 2002). Salinity is a common environmental challenge all around the world. Soil salinity significantly reduces the productivity of economical important plants. Salts in the soil water prevent the growth of plants because of two reasons. First, the presence of salt in the soil solution reduces the ability of the plant to absorb water, and consequently growth rate will be decreased. This is called the osmotic or water-deficit effect of salinity. Secondly, if excessive amounts of salt enter the plant in the transpiration stream, the cells can be damaged in the transpiring leaves and this may cause more reductions in growth. This is called the salt-specific or ion-excess effect of salinity (Yao & Fang, 2009). In general, seed germination is considered as an important stage in plants growth cycle. In fact, it controls the beginning of seedling growth. Salinity factor’s has a critical influence on the seed germination which is the most sensitive step in plant growth (Ahmad et al., 2009) and the seeds exposed to unfavorable conditions like salinity may have to compromise the seedlings development (Albuquerque & Carvalho, 2003). Tomato (Solanum lycopersicum) belongs to the family Solanaceae that includes several additional economically important crops such as potato, pepper and eggplant. Tomato is native to Southern North America, Central and South and it is the most popular and second important frequently consumed vegetable crop grown throughout the world after potato (Dorais et al., 2008). It is recognized as a highly valuable and nutritious food. These days, tomato is one of the major vegetable throughout the world. It is grown both in the open and green house. Moreover, it is fruit contain a number of health-beneficial

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compounds, such as high potassium content, iron, vitamins A, B, C and lycopene (Preedy, 2008). 1.2 Problem Statements and Objectives of Study The main aim of this work is to study the effects of salt stress on tomato plant. Many studies had been conducted regarding the effects of salt stress towards growth, physiology and productivity of tomato plant but study on the effect of salinity on seed germination and antioxidant activity of tomato plant in response to salt stress is still limited. Thus, this present research was conducted because seed germination is a critical stage in the life cycle of plants and salt tolerance during germination is crucial for the establishment of plants that grow in saline soils. In addition, study on the effects of NaCl on the germination and antioxidant activity has never been conducted of tomato MT1. Salt stress study also contributes to productivity of agriculture and ways to overcome the problem of salinity in agriculture. This is important to increase food production and fulfill food demand worldwide. The objectives of study are:

1. To study the effects of different concentrations of NaCl on the germination and early seedling growth of tomato.

2. To study the effects of different concentrations of NaCl on the histological leaves of tomato seedlings

3. To study the effects of different concentrations of NaCl on anti- oxidative profiling of tomato plant

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